Porcupine in the Late Neogene and Quaternary of Georgia

saqarTvelos mecnierebaTa erovnuli akademiis moambe , t. 4, #3, 2010BULLETIN OF THE GEORGIAN NATIONAL ACADEMY OF SCIENCES, vol. 4, no. 3, 2010

© 2010 Bull. Georg. Natl. Acad. Sci.

Palaeobiology

Porcupine in the Late Neogene and Quaternary of Georgia

Abesalom Vekua*, Oleg Bendukidze**, Maia Bukhsianidze**,Nikoloz Vanishvili**, Jordi Augusti†, Bienvenido Martinez-Navarro§,Lorenzo Rook§§

* Member of the Georgian Academy of Sciences, Institute of Paleobiology, Georgian National Museum** Institute of Paleobiology, Georgian National Museum† ICREA, IPHES, Universidad Rovira Virgili, Tarragona, Spain§ Museo de Paleontologia Orce, Spain§§ Dipartiménto di Sciènze della Terra, Università di Firenze, Firenze, Italy

ABSTRACT. The porcupine family (Hystricidae) is notable for the diversity of fossil and modern forms. Due tovagueness of morphological characters their taxonomy is not yet established. Lately an interesting work by vanWeers and Rook was published about the taxonomy of European, Asian and African porcupines in which the authorspropose relatively natural and stratigraphically reasonable views. © 2010 Bull. Georg. Natl. Acad. Sci.

Key words: porcupine, Pliocene, Paleolithic, Neolithic.

Introduction.The porcupine history in Eurasia starts in the Late

Miocene. No earlier remains are found yet. However, forsome unspecified data Oligocene age is also proposed.Authentic relicts of porcupines were found in Vallesian(MN10) and Early Turolian (MN11). Smaller forms areattributed to the species Hystrix parvae (Kretzoi) andlarger and advanced ones to H. primigenia. The latterforms are known from the Neogene of Eastern Europe.Stratigraphically H. primigenia occurs in the Early andMiddle Turolian (MN11/12) and persists nearly to theend of Ruscinian (MN15). It should be noted that relictsof a porcupine (Hystricinae) [1], found in the Maeotiansediments of David Gareji desert (Eastern Georgia), mostprobably belong to a species of Hystrix because no otherHystricidae ever inhabited the South Caucasus. Unfortu-nately, porcupine fossils found in the David Garedji desertwere lost and now it is impossible to revise the material.

The Hystricidae family is divided into two subfa-milies: relatively primitive, long-tailed Atherurinae Lyon(1907) and true porcupines - Hystricinae Murray, 1866.

The latter subfamily comprises several modern and fossilforms. In this article we mainly consider species of thegenus Hystrix from Dmanisi.

Van Weers and Rook [2] significantly simplifiedporcupunes’ taxonomy and synonymised morphologi-cally and stratigrafically similar forms, which undou-btedly will facilitate thorough study of the group underconsideration.

Masini and Rook [3] carried out a serious revisionof the described porcupines from the reference Europeanlocalities (Pikermi, Wenze, Perpignan, Etouaires, Perrier)and identified morphological characters specific to theconsidered forms. Later, Sen [4], considering theaforementioned morphological characters, attributed aporcupine from Perpignan as a new species - Hystrixdepereti. This form obviously differs from typical species- H. primigenia by relatively high crowned teeth andmassive incisors, while a larger porcupine from Etouairesis attributed to another species - Hystrix refossa Gerv.Thus, in the Late Neogene three species are present inthe following stratigraphic order: H. primigenia, H.depereti and H. refossa; nevertheless phylogenetic

Porcupine in the Late Neogene and Quaternary of Georgia 141

Bull. Georg. Natl. Acad. Sci., vol. 4, no. 3, 2010

relations between them are not yet clear.Large fossil porcupines easily split into two groups:

1) forms with comparatively brachyodont teeth ofMiocene and Pliocene age and 2) porcupines havingnearly as hypsodont teeth as the representatives ofmodern species of the genus Hystrix. The mentioneddivision is based on tooth crown height to teeth grindingsurface length ratio [2].

The Dmanisi porcupine is undoubtedly similar tothe second group by having relatively large andhypsodont teeth. Phylogenetic kinship with the afore-mentioned forms is also confirmed by the presence offour roots on M1 and M2 (unlike H. cristata. In thisspecies the same teeth have three roots). Presumably,the Dmanisi porcupine may be considered as an ancestorof a modern H. indica. The Dmanisi form has larger teeth,yet the difference in size between Dmanisi porcupineand H. indica is negligible.

Tchernov [5] described porcupines from Ubeidja andattributed them to H. indica, stating that the Ubeidijaporcupine is morphologically identical to H. angressiapart from having relatively small dentition. At the sametime Weers and Rook synonymise H. angressi with H.refossa. Several European and Western Asian speciesas well - H .major, H. etruscus, H. angressi and SouthAfrican H. makapensis are synonymised by thementioned authors with H. refossa as well [2]. Weconsider that the Dmanisi porcupine and the one foundin the Mousterian layers of Tsutskhvati cave [6] shouldbe attributed to H. refossa.

Hysrtix refossa Gervias, 1852

Figs.1-4

Synonymy: H. major Gervias, 1859; H. etruscaBosco, 1898; H. angressi Frenkeli, 1970; H. makapensisGreenwood, 1958; Hystrix sp. Vekua, 1978.

Site: Dmanisi, end of Villafranchian, MN17.Fossil material: D2718 - a fragment of maxillary bone

with complete tooth rows on both sides (P2-M3); D4120- upper left, very worn, isolated M1; D4140 - isolated, alittle alveolar with broken end, upper incisor; D4509 -nearly complete mandible with ascending branch on bothsides, with incisors and P4-M3, on the left hemimandibleand with incisor and P4 – M2 on the right one; D3878 -fragment of a right hemimandible with moderately warnP4 and very worn M1.

Description and comparison:The Dmanisi Hystrix refossa is a comparatively large

rodent with teeth measurements nearly similar to H.primigenia and H. depereti. However there aredifferences in morphology. The Dmanisi porcupine hassufficiently hypsodont teeth; ratio of crown height tolength of occlusal surface of its teeth always exceeds100. The same ratio is characteristic of the teeth of H.refossa and also of the representatives of modern Hystrixsubspecies.

The Palate of the Dmanisi porcupine is longitudinallyslightly concave in the middle in the form of a narrowgroove. Anterior edge of the choana is rounded andreaches the middle of M3. On maxilla tooth rows are notparallel to each other but slightly broadened mesially(Fig.1).

Fig. 1. Dmanisi Hystrix (Hystrix) refossa Gerv., maxilla,occlusal view.

P4 is weakly worn, it is obliquely inserted in maxillaand inclined backwards towards other teeth. All the fourroots are weak, isolated, with open pulp. Lingualhypoflexion is deep and joins the labial paraflexion andthus the anterior part is separated. At the labial edge ofthe tooth three short folds are present.

M1 and M2 have nearly the same size, somewhatelongated, moderately worn, labially with three folds.

Fig. 2. Dmanisi Hystrix (Hystrix) refossa Gerv., M1 dex, occlusalview.

142 Abesalom Vekua, Oleg Bendukidze, Maia Bukhsianidze, Nikoloz Vanishvili


Distal crescent like folds are longer than others. Thereare three isolated knots on M1 and two on M2; M3 israther similar to the previous molars.

The upper incisor (D4140) is represented only bythe anterior half. Occlusal surface has the shape ofelongated rectangle, much curved arclike. Yellow enamelcovers only the upper part of the tooth and expands abit to the sides.

The dimensions of the incisor are: width - 6.7 mm,antero-posterior diameter – 8 mm. It should be remarkedthat in H. refossa upper incisor width is 5.6 mm andantero-posterior diameter – 7 mm. Obviously, the Dmanisiporcupine has somewhat stronger and massive upperincisors.

Almost complete mandible (D4509) of porcupine ispresent in the Dmanisi collection. There are incisor, P4,M1 and M2, on the right, while in the left incisor, P4,M1, M2, and M3. The mandible is rather massive (theheight of the horizontal branch at M1 level reaches 27mm, the width at the same point is 22.4 mm), rather largeforamen mentale is remarkable on the mandible (DAP is4.3, the height – 3.2 mm). P4 is not yet worn, while molarsare moderately worn. Lower incisors, in comparison withthe upper ones, are less curved, they are rather robust.Enamel covers the anterior part and expands on the sides.Dimensions of the incisor are: length – 92 mm, DVL - 7.4mm, DAP – 8.5 mm.

P4 has the shape of an elongated cylinder, crown ishigh enough. A deep fold is developed labially, the grooveof which reaches the tooth root. Lingually four morefolds are observable, one of them - a mesofacetidpenetrates deeply into the occulsal surface. Root pulpsare open.

M1 and M2 have similar morphology: labially withwell developed hypofossetid and lingually with similarlydeveloped mesofossetid. There are four isolated knotson the occlusal surface of the tooth. A peculiar characterof M1 is the existence of an isolated enamel column onthe posterior wall of the labial fold. Similar column isobserved on the left M1 of the same jaw as well.Occurrence of the mentioned additional column is rarein porcupines but Hytrix major, which is considered tobe a synonym of H. refossa, has it on M3 [7]. It is possiblethat this character is peculiar to H. refossa.

General size and morphology (folds, fossetids) ofDmanisi lower dentition is similar to H. angress, whichis also considered as a synonym of H. refossa [5].According to our observations the Dmanisi porcupinediffers from the Ubeidija porcupines only by the presenceof more fossets and fossetids, which may be explainedby their intense wear.

Dmanisi H. refossa differs from the typical H.primigenia by having somewhat smaller teeth, and moresignificantly by difference in the development of foldsand distribution of elements on the occlusal surface.The Dmanisi porcupine has deep lingual and labial foldsthat extend vertically along the whole length of thecrown, reaching roots. The mentioned folds areobservable even on very strongly worn teeth.

Although it has been demonstrated [8,9] that theocclusal patterrns may show a great variabilitydepending on the stage of wear of the tooth, in thecase of H. primigenia these folds are relatively lessdeveloped, so that on moderately worn teeth their tracesare weak and on intensively worn off teeth theycompletely disappear; the presence of an additionalcolumn in the Dmanisi porcupine is another remarkabledifference from the Pikermi form as well.

Modern H. cristata and fossil H. subcristata differfrom the Dmanisi form by smaller dimensions, lessernumber of closed knots and absence of isolated columns.

The Dmanisi porcupine differs from modern H.hirsutirostris by larger teeth and quantity of roots (four

Fig. 3. Dmanisi Hystrix (Hystrix) refossa Gerv., mandible,lateral view.

Fig. 4. Dmanisi Hystrix (Hystrix) refossa Gerv., mandible,occlusal view.



and three roots respectively). Researchers consider thischaracter to be diagnostic.

Some years ago Sharapov synonymised H. trofimovito H. primigenia [10]. This idea was not shared by N.Shevyreva, who defined the Kuruksai porcupine as anew species – H. trofimovi [11]. Later Weers and Rooksynonymised it with H. primigenia [2]. In any case thefossil porcupine from Kuruksai is different from theDmanisi form by having three roots (four roots inDmanisi) and missing of postmetaflexes.

The Dmanisi porcupine significantly differs in sizefrom the Binagady H. vinogradovi (the latter is smaller)– a species defined by Argyropulo in 1941 fromBinagady (Middle Pleistocene). However, he did notprovide any description of the fossil [12], only mention-ing that the Binagady porcupine is the smallest amongmodern forms. A Comparatively vast description of theBinagady fossil porcupine is given by I. Gromov [13].

Somewhat later porcupine fossils were found in theAcheulian-Mousterian cultural layers of the Kudarocave. According to Baryshnikov and Baranova, theKudaro porcupine is a subspecies of the Binagadyporcupine and is defined as H. vinogradovi kudarensis[14], yet Weers [15] attributes the Binagady porcupineto the subspecies Acanthion [15].

Study of the Dmanisi porcupine naturally involvedrevision of available Quaternary porcupine material fromWestern Georgia. The major part of the fossils comesfrom the excavations of archaeological sites. Relativelygood material is found in Tetri Mghvime (Tskaltuboregion, Upper Paleolithic, Eneolithic). The very fact thatporcupine remains are found in the Eneolithic strata isof great importance, because it indicates that this rodentinhabited Western Georgia during the Holocene and wasa rare element of Quaternary fauna.

Mainly small-sized porcupines inhabited Western

Georgia during the Pleistocene and Holocene. It doesnot differ essentially from the Binagady and Kudaroforms in size and morphology.

It must be also noted that [15] united small-sizedporcupines, found in the Southern Caucasus and CentralEurope, fall into the subgenus Acanthion. It is clear thatH. vinogradovi from Binagady and H. vinogradovikudarensis from Kudaro fall into this subgenus.

As already remarked above, the porcupines ofWestern Georgia do not differ essentially in their sizeand morphological features of teeth from the small-sizedporcupines of Kudaro, Binagady and Europe, that areunited into the subgenus Acanthion of the genusHystrix. That is why we attribute porcupines found inthe Pleistocene-Holocene sediments of Western Georgiato the subgenus Acanthion without hesitation.

Below we give a brief description of porcupine teethfrom Tetri Mghvime:

Hystrix (Acanthion) vinogradovi Argyropulo, 1941Synonyms (according to Weers):H. shaubi Brunner, 1954H. cristata minor Malez, 1963H. vinogradovi atavus Janossy, 1972H. vinogradovi kudarensis Baryshnikov, Baranova

1982.Material: Fragment of maxilla (1175, sin., Eneolithic)

with rather worn M1/-M3/; fragment of maxilla (916, dex.,Paleolithic) with only one almost unworn (P4/) tooth;mandible (1287, sin., Eneolithic) with complete tooth row(P/4-M/3); fragment of mandible (1355, sin., Eneolithic)with slightly worn M/2; upper incisor (520, sin.,Eneolithic); upper incisor (800, Paleolithic). All materialcomes from the Upper Paleolithic and Eneolithic strataof Tetri Mghvime, together with fossils of the bear andother animals.

There is nearly nothing to say about porcupinemaxilla from Tetri Mghvime because of its very fragmen-tary nature. Only one fragment of maxilla (1175) is of

Fig. 5. Tetri Mghvime. Hystrix (Acanthion) vinogradovi,maxilla (M1-M3, sin. Eneolithic), occlusal wiev.

Fig. 6. Tetri Mghvime. Hystrix (Acanthion) vinogradovi,maxilla (P4, dex. Paleolithic), occlusal view.



interest, on which three well-preserved molars (M1/-M3/)are preserved; they remind us a much diminished copyof the upper jaw and teeth of a large-sized porcupine.

Actually all teeth of porcupine are present in TetriMghvime: isolated upper incisor, P4/ (N916) attached toa small fragment of maxilla and all three molars in thesecond upper jaw (1175).

P4/ is almost unworn. On the occlusal surfacehypoflexia and mesoflexia are transversally connectedand protruded forward in the form of an obtuse angle.In the mesial and distal parts of tooth round form fossetsare located. Lingual groove (hypoflexia) is well visibleon the upper part of the crown; however, the verticalgroove disappears towards the root. We should notehere that the small porcupine from Western Georgia hasfour roots, of which medial roots are weak and separated,and buccal roots are united into one massive root.Hypoflexia is obliquely protruded forward a little andconnected with mesoflexia, so that the distal part of thetooth seems separated as an independent cusp.

Molars are placed close to one another. Teeth aresmall and almost equal in size. Signs of two meso- andhypoflexia are notable on the buccal wall of M1/ andM2/.

Mandible and lower teeth. An almost completemandible with tooth row (P/4-M/3) is found in TetriMghvime, as well as a fragment of mandible with weaklyworn M/2.

The mandible of H. (Acanthion) vinogradovi fromTetri Mghvime has somewhat different morphologicalfeatures from other porcupines, especially from H. indica.First of all there is a size difference. The porcupine fromTetri Mghvime is much smaller, besides it differs byreduced robustness, by location of joint surface andforamen mentale, by the height and length of theascending branch and other features.

Lower teeth are in mid wear stage and have adiagonally elongated shape. On the occlusal surface all

fossets are looped. Shape and number of loops on teethare different. P/4 displays three knotted fossets, M/1 -two, M/2 - three and M/3 – five.

General remarksIt is noteworthy that in Eastern Georgia there is no

occurrence of porcupine during the entire Pleistoceneand Holocene regardless the fact that there are a lot ofpaleontological and archaeological sites of this age. Thesituation is opposite in Western Georgia - porcupineremains are common during the Pleistocene and persistuntil the very end of the Holocene.

According to Pidoplichko [16], the porcupine is aninhabitant of xerophyte landscape, where snow cover isnegligible or nonexistent. In the Pliocene-Holocenexerophyte landscape was well developed in EasternGeorgia but in Western Georgia such landscapesoccupied a limited area.

Some scientists consider that intense growth offorests in Western Georgia caused extinction of porcu-pines in the Holocene. This assumption seems uncon-vincing. Western Georgia never suffered from the lackof vast regions covered with forests. It is more accept-able to consider that the prehistoric man of the Earlyand Late Stone Age often used porcupines for subsisten-ce and intensively hunted them as they are easy to catch;finally this caused the extinction of the rodent.

Baryshnikov’s and Baranova’s [14] view on theextinction of porcupines in Western Georgia is worthyof attention. The mentioned authors suppose that theextinction of porcupines in Western Georgia at the endof the Pleistocene was related to glaciation and generalfall of temperature. Unfortunately, the authors neglectthe facts of existence of porcupines’ remains in TetriMghvime (Eneolithic, Neolithic), Dzudzuana (Eneolithic)and Tsona (Mesolithic, Eneolithic).

It must be noted that the great glaciation in Russiadid not cause essential changes in the composition offauna of West Georgia. They only mention the verticaldown shift of the lower border of snow and glacier bysome hundred meters invoked displacement of mountainforms to valleys, which is generally expectable.

Let us recollect that Georgian territory was a peculiarrefugium where warm and moderately humid climate,mosaic landscape and favorable living conditions werepreserved during the Plio-Pleistocene and later. Thisrefugium conditioned the preservation of relicts of earlierflora and fauna. Here are some facts. The latestanthropoid ape in Eurasia (Dryopithecus garedziensis,[17] was found in Gareji desert (Maeotian sediments); Inthe former Soviet Union territory the genus Macaca in

Fig. 7. Tetri Mghvime. Hystrix (Acanthion) vinogradovi,mandible (P4-M3 sin. Eneolithic), lateral view.



the Pleistocene is known only from the Kudaro cave[18]; In Central and Eastern Europe Ursus spelaeus wascompletely extinct by the end of the Pleistocene andonly in Western Georgia (Abkhazia) it is often found inMesolithic cultural strata [19]; Woodchuck (Marmotasp.) does not inhabit Georgia at present, but in thePleistocene (Zemo Orozmani) and Holocene (Kutaisienvirons) it was probably widely spread and in spite ofgreat glaciation in Russia hamster’s populations surviveduntil the historical period in Western Georgia [20].

In South Caucasica, in Azerbaijan and Armenia, the

porcupine is considered to be a modern faunal element,while in Georgia, some scientists practically exclude thepossibility of its occurrence in modern fauna.

V. Shidlovski in his monograph “Identification guideof South Caucasus rodents” [21] does not even mentionthe porcupine in the list of Georgian fauna. Meanwhile,it has turned out that this rodent is present in Eastern

Fig. 8. Tetri Mghvime. Hystrix (Acanthion) vinogradovi,mandible (P4-M3 sin. Eneolithic), occlusal view.

Fig. 9. Tetri Mghvime. Hystrix (Acanthion) vinogradovi,mandible (M2 sin. Eneolithic), lateral view.

Table 1

Measurements of the upper teeth of hystrix

H. r

efos

sa D

man

isi

H. p

rimig

enia

Pi

kerm

i

H. d

eper

eti

Perp

igna

n

H. i

ndic

a U

beid

iya

H. a

ngre

ssi

Rece

nt

H. i

ndic

a R

ecen

t

H. v

inog

rado

vi

kuda

rens

is

H. v

inog

rado

vi

Arg

. Bi

naga

di

H.(A

cant

hion

) vi

nogr

adov

i Te

tri M

ghvi

me

(G

eorg

ia)

Weers, Rook 2003

Ternov, 1986 Baryshnikov, Baranova, 1982

Diameter I 8-8.5 7.0 - 8.3 7.0 - - - 4.9

P4 – M3 38 39.3

Lengh

P4

Width

9.0

8.0

10.8

11.2

11.2-11.3

9.9-10.3

10.5

9.6

9.0

8.0

Length

M1

Width

9.0

8.4

9.0

10.2

9.6-10.8

8.0-10.7

9.0

7.7

7.3

7.7

9.4

7.7

6.8-7.0

5.5-6.3

6.5-6.7

5.5-5.7

5.6

8.0

Length

M2

Width

9.0

8.4

9.8

10.8

9.5

8.2

8.0

7.4

5.6

6.4

Length

M3

Width

8.7

7.4

9.0

9.4

8.7

9.0

9.1

7.6

7.5

5.9

7.7

6.5

6.5+

5.5

7.3

6.2



Georgia (Vashlovani reserve, Kakheti - the territoryadjacent to Azerbaijan) [22]. Besides, shepherds andemployees of the protected territories often came acrossporcupines on the mentioned territory. Attention shouldbe paid to rather frequent cases of finding porcupinequills in the Iori river valley. We also found porcupinequills and brought them to Tbilisi as a confirmation ofporcupine presence on the territory of Georgia.

As we already mentioned above, some researchersexclude the possibility of porcupine presence on theterritory of Georgia. We are far from thinking that theaforementioned researchers stated their ideas withoutverification of facts. It is more reasonable to assumethat porcupines entered Eastern Georgia very recentlyand reproduced rapidly due to favorable xerophytelandscape and warm climate

In Western Georgia the case is somewhat different.No porcupines are observed in modern fauna but theywere abundant during the Pleistocene-Holocene. Remainsof porcupines in Western Georgia are mainly representedby leftover bones in cultural layers of Paleolithic andNeolithic sites. At present in Western Georgia porcupine

remains have been found at the following cave sites:Tetri Mghvime (Neolithic) [23], Okumi (Lower Paleolithic)[24] Tsona (Mousterian) [25] Ortvala Mghvime(Mousterian, Mesolithic, Eneolithic) [26], Djruchula(Upper Paleolithic) [23] Dzudzuana (Eneolithic) [23],Sakajia (Mousterian) [23] Kudaro (Acheulian,Mousterian) [14;27]. It should be noted that amongleftover bones porcupine is represented by few remainsand researchers attribute them either to various speciesor do not determine them at all. As stated above on theterritory of Eastern Georgia no porcupine remains werefound from Quaternary times.

In 1972, at Kvabebi site (near Sighnaghi, Kakhetiregion, Eastern Georgia) unique fauna of the MiddleAkchagylian age contained two isolated teeth that werepresumably attributed to H. cf. primigenia [28, 29]N. Shevyreva [11] did not share Vekua’s determinationand assumed that the Kvabebi porcupine was closer toH. cristata. We compared the teeth of the Kvabebiporcupine with those of H. primigenia and otherPliocene forms once more and assumed that with thesetwo isolated and intensively worn off teeth it is

Table 2

Measurements of the lower teeth of hystrix

H

. ref

ossa

Dm

anis

i

H. p

rimig

enia

Pi

kerm

i

H. d

eper

eti

Perp

igna

n

H. i

ndic

a U

beid

ija

H. a

ngre

ssi

Rece

nt

H. i

ndic

a R

ecen

t

H. v

inog

rado

vi

Kud

aren

sis

H. v

inog

rado

vi

Arg

. B

inag

ady

H.(A

cant

hion

) vi

nogr

adov

i Te

tri M

ghvi

me

(Geo

rgia

)

Weers, Rook 2003

Thernov, 1986

Baryshnikov, Baranova, 1982

Diameter I 8.4 - - 7.2 7.4 6.3 4.8-4.9 - -

P4 – M3 42 41.6-43.4 47.1 _ _ _ 29. 7 28.1 24-27

Lengh

P4

Width

12-13

6.7-9.5

11.2-11.3

9.9-10.3

12.3-13.2

10-11.8

8.5

7.3

10.8

8.2

9.0

6.7

8.0

6.1

Length

M1

Width

10.7-11

7.8-9.5

9.0-11.6

8.4-9.4

11.1-13.1

8.9-10.8

9.2

8.0

9.0

7.5

7.2

6.8

7.7-8.2

6.3-6.9

6.9-8.5

5.0-6.5

5.3

5.5

Length

M2

Width

11.2-11.6

7-7.6

-

-

-

-

10.1

8.3

9.6

7.3

8.2

7.0

-

-

-

-

6.3

6.0

Length

M3

Width

9.3

6.6

9.4-9.8

8.4-8.6

9.6-10.5

8.4-9.0

9.6

7.2

10

7.4

8.1

6.4

6.0-7.3

5.0-5.6

5.8-6.7

4.5-5.6

6.2

6.4



impossible to give an exact determination of the speciesof the Kvabebi porcupine. Nevertheless, it should beremarked that the Kvabebi porcupine is more similar toH. primigenia than to H. cristata according to size and

Fig. 10. Lower and Upper dentition (P4/, M1-2/; P/4, M/1-2) graphs showing the range of occlusal length (left column) andhypsodonty index (right column) for extant Hystrix cristata, hypsodontic Plio-Pleistocene species (Hystrix refossa) andbrachydont species (Hystrix primigenia, Hystrix depereti). Hystrix specimens from Dmanisi fall in the range of Hystrix refossa.Comparative data from [2,9,15].

morphology of teeth. Besides, we would like to mentionthat the geological age of the Kvabebi fauna (MN16)[30] does not exceed much the upper boundary ofstratigraphic distribution of H. primigenia [2].



paleobiologia

maCvzRarba saqarTvelos gvian neogenur da meoTxeulfaunaSi

a. vekua*, o. benduqiZe,** m. buxsianiZe,** n. vaniSvili**, J. agusti†,b. martines-navaro§, l. ruki§§

* akademiis wevri, saqarTvelos erovnuli muzeumi, paleobiologiis instituti, Tbilisi** saqarTvelos erovnuli muzeumi, paleobiologiis instituti, Tbilisi

† adamianis paleobiologiis instituti, taragona, espaneTi§ paleontologiis muzeumi, orse, espaneTi§§ florenciis universiteti, mecnierebaTa departamenti, florencia, italia

statia eZRvneba saqarTvelos teritoriaze paleocensa da meoTxeulSi maCvzRarbebis gavrce-lebas. aRmosavleT saqarTveloSi maCvzRarbebis (Hystrix) namarxi naSTebi ar gvxvdeba. dasavleTsaqarTveloSi pliocen-meoTxeulis manZilze maCvzRarba farTod yofila gavrcelebuli, rac uaxlesipaleontologiuri aRmoCenebiT dasturdeba.

saqarTveloSi Tanamedrove faunaSi maCvzRarbebis arsebobas specialistebi uaryofen. sinamdvi-leSi ki azerbaijanis momijnave kaxeTis teritoriaze maCvzRarba dResac binadrobs, rac dasturdebabuxnikaSvilisa da CxikvaZis monacemebiT (2004). savaraudoa, rom maCvzRarba azerbaijanidan gadmovidakaxeTis teritoriaze da aq xelsayreli saarsebo pirobebis gamo damkvidrda.

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Received July, 2010

Porcupine in the Late Neogene and Quaternary of Georgia

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